Rhacophorus Kuhl and Van Hasselt, 1822

Class: Amphibia > Order: Anura > Family: Rhacophoridae > Subfamily: Rhacophorinae > Genus: Rhacophorus
45 species

Rhacophorus Kuhl and Van Hasselt, 1822, Algemeene Konst-en Letter-Bode, 7: 104. Type species: Rhacophorus moschatus Kuhl and van Hasselt, 1822 (= Hyla reinwardtii Schlegel, 1840), by monotypy (see comments by Dubois, 1989 "1988", Alytes, 7: 101–104, and Ohler and Dubois, 2006, Alytes, 23: 123–132).

RacophorusSchlegel, 1826, Bull. Sci. Nat. Geol., Paris, Ser. 2, 9: 239. Incorrect subsequent spelling.

RhacoforusPalacky, 1898, Verh. Zool. Bot. Ges. Wien, 48: 374. Incorrect subsequent spelling.

Huangixalus Fei, Ye, and Jiang, 2012, Colored Atlas Chinese Amph. Distr.: 598. Type species: Rhacophorus translineatus Wu, 1977, by original designation. Provisionally retained in this synonymy because its recognition would render Rhacophorus paraphyletic (DRF). 

Vampyrius Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 262. Type species: Rhacophorus vampyrus Rowley, Le, Thi, Stuart, and Hoang, 2010. See comment under Rhacophorus vampyrus. Synonymy by Mahony, Kamei, Brown, and Chan, 2024, Vert. Zool., Senckenberg, 74: 253. 

English Names

Flying Frogs (Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 113).

Parachuting Frogs (Harvey, Pemberton, and Smith, 2002, Herpetol. Monogr., 16: 46).

Tree Frogs (Dinesh, Radhakrishnan, Deepak, and Kulkarni, 2023, Fauna India Checklist, vers. 5.0 : 14).


Southern India to Bhutan and eastern Xizang (China) east and south to Hunan, Hainan, Yunnan, and Guangxi, China, though Myanmar, Thailand, Laos, Cambodia, and Vietnam to Sumatra, Borneo, and Sulawesi (Indonesia) and Philippines. 


Dubois, 1982, Bull. Mus. Natl. Hist. Nat. Paris, Sect. A, Zool., 4: 271–276, discussed the nomenclature of the type species. Inger, 1954, Fieldiana, Zool., 33: 370–393, reviewed the Philippine species. Taylor, 1962, Univ. Kansas Sci. Bull., 43: 480–505, reviewed the species of Thailand. Inger, 1966, Fieldiana, Zool., 52: 279–340, reviewed the Bornean species. Bourret, 1942, Batr. Indochine: 408–449, reviewed the southeastern Asian species. Liu and Hu, 1961, Tailless Amph. China: 272–273, reviewed the Chinese species. Dubois, 1987 "1986", Alytes, 5: 75–85, discussed the subgenera and species groups discussed in the accounts (but see Polypedates). Brown and Alcala, 1994, Proc. California Acad. Sci., Ser. 4, 48: 206, provided a key to the species of the Philippines. See comment under Polypedates. Harvey, Pemberton, and Smith, 2002, Herpetol. Monogr., 16: 46–92, discussed the systematics of the species from Sumatra and Java, rejected the species groups of Rhacophorus formulated by Dubois, 1987 "1986", Alytes, 5: 7–95, as undiagnosed and criticized Leptomantis as constructed by Dubois, 1987 "1986", Alytes, 5: 7–95, and Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 305–352, as likely polyphyletic. Malkmus, Manthey, Vogel, Hoffmann, and Kosuch, 2002, Amph. Rept. Mount Kinabalu: 199, provided a key for the species of Borneo. Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006, Bull. Am. Mus. Nat. Hist., 297: 139, considered Rhacophorus to be the sister taxon of Polypedates + Chiromantis. Matsui and Panha, 2006, Zool. Sci., Tokyo, 23: 477–481, commented on the validity of Polypedates and Leptomantis compared with Rhacophorus. Grosjean, Delorme, Dubois, and Ohler, 2008, J. Zool. Syst. Evol. Res., 46: 169–176, and Li, Che, Bain, Zhao, and Zhang, 2008, Mol. Phylogenet. Evol., 48: 302–312, provided evidence that Rhacophorus is composed of two major clades. Unfortunately, at present the assignment of the available names, Rhacophorus and Leptomantis is unclear. Orlov, Nguyen, and Ho, 2008, Russ. J. Herpetol., 15: 67–84, provided a key to the species of Vietnam. Fei, Hu, Ye, and Huang, 2009, Fauna Sinica, Amph. 2: 766–868, provided a key to and accounts for the species of China, and recognized species groups. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, provided a tree of examplar species and confirmed the placement of a monophyletic Rhacophorus as the sister taxon of Feihyla + Polypedates­ + Taruga (which they considered part of Polypedates). Li, Liu, Chen, Wu, Murphy, Zhao, Wang, and Zhang, 2012, Zool. J. Linn. Soc., 165: 143–162, reported on the molecular phylogenetics of the Rhacophorus dugritei group. Li, Li, Murphy, Rao, and Zhang, 2012, Zool. Scripta, 41: 557–570, reported on the systematics of Rhacophorus, delimiting three major subgeneric taxa, which they declined to name pending additional work—Clade A (which likely carries the name Leptomantis): Rhacophorus angulirostris, Rhacophorus belalongensis, Rhacophorus bimaculatus, Rhacophorus fasciatus, Rhacophorus feae, Rhacophorus harrissoni, Rhacophorus monticola, Rhacophorus penanorum, and Rhacophorus rufipes; Clade B (which carries the name Rhacophorus): Rhacophorus annamensis, Rhacophorus bipunctatus, Rhacophorus calcaenus, Rhacophorus chuyansinensis, Rhacophorus kio, Rhacophorus lateralis, Rhacophorus malabaricus, Rhacophorus nigropalmatus, Rhacophorus orlovi, Rhacophorus pardalis, Rhacophorus reinwardtii, Rhacophorus rhodopus, Rhacophorus translineatus, and Rhacophorus verrucopus; Clade C (unnamed): Rhacophorus arboreus, Rhacophorus burmanus, Rhacophorus chenfui, Rhacophorus dennysi, Rhacophorus dorsoviridis, Rhacophorus duboisi, Rhacophorus dugritei, Rhacophorus dulitensis, Rhacophorus feae, Rhacophorus hui, Rhacophorus hungfuensis, Rhacophorus maximus, Rhacophorus minimus, Rhacophorus moltrechti, Rhacophorus nigropunctatus, Rhacophorus omeimontis, Rhacophorus puerensis, and Rhacophorus schlegeliiLi, Li, Klaus, Rao, Hillis, and Zhang, 2013, Proc. Natl. Acad. Sci. USA, 110: 34413446, confirmed that Rhacophorus is the sister taxon of Polypedates + Taruga + Ghatixalus + Feihyla and also confirmed the deep structure within Rhacophorus noted by Li et al., 2012. Matsui, Shimada, and Sudin, 2013, Curr. Herpetol., Kyoto, 32: 112–124, provided evidence regarding the molecular phylogenetics of the Rhacophorus reinwardtii group.  Ostroshabov, Orlov, and Nguyen, 2013, Russ. J. Herpetol., 20: 301–324, discussed the species similar to Rhacophorus hoanglienensis and Rhacophorus orlovi in northern Vietnam, with the naming to two species. Nguyen, Matsui, Eto, and Orlov, 2014, Russ. J. Herpetol., 21: 274–280, discussed a molecular tree of the species found in Vietnam, which, based on a small amount of evidence and inherently large amounts of missing terminals, did provide evidence of unnamed species and previous unsuspected relationships. Mo, Chen, Liao, and Zhou, 2016, Asian Herpetol. Res., 7: 139–150, provided a Bayesian tree based on mtDNA data for the genus. Pan, Zhang, Wang, Wu, Kang, Qian, Chen, Rao, Jiang, and Zhang, 2017, PeerJ, 5(e3995): 1–22, reported on the molecular biogeography of the genus. O'Connell, Smart, Smith, Hamidy, Kurniawan, and Fujita, 2018, J. Biogeograph., 45: 929–940, reported on the molecular phylogenetics and biogeography of the species focusing on the biogeography of the Sunda Shelf. Jiang, Jiang, Ren, Wu, and Li, 2019, Asian Herpetol. Res., 10: 1–12, partitioned nominal Rhacophorus into three genera in the relationship Rhacophorus + (Leptomantis + Zhangixalus), rendering Rhacophorus a much smaller genus with a much restricted geographic range. Literature of the species of Rhacophorus (sensu lato) prior to this publication therefore may apply also to species of Leptomantis and Zhangixalus.  Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 427, recovered Vampyrius far from Rhacophorus and the sister taxon of Gracixalus (their Gracixalus + Orixalus); this was rejected by the evidence of Chan, Hutter, Wood, Grismer, and Brown, 2020, Proc. R. Soc. London, Ser. B, Biol. Sci., 287 (20202102): 1–2, as discussed by Mahony, Kamei, Brown, and Chan, 2024, Vert. Zool., Senckenberg, 74: 253. Tang, Xiao, Liu, Wang, Yu, and Du, 2024, Zoosyst. Evol., 100: 625–643, reported on the molecular phylogenetics of the Rhacophorus rhodopus and Rhacophorus bipunctatus complexes. 

Contained taxa (45 sp.):

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